gene list for cucurbita species, 2004 - cucurbit...

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Gene List for Cucurbita species, 2004 Harry S. Paris A.R.O., Newe Ya’ar Research Center, Ramat Yishay 30-095 (Israel) Rebecca Nelson Brown Oregon State University, Corvallis, OR 97331 (U.S.A.) A complete list of genes for Cucurbita species was last published 12 years ago (33). Since then, only updates have been published (72, 73). The genus Cucurbita L. contains 12 or 13 species (50). As far as is known, all have a complement of 20 pairs of chromosomes (2n = 40). This new gene list for Cucurbita contains much more detail concerning the sources of information, being modeled after the one for cucumber presented by Wehner and Staub (96) and its update by Xie and Wehner (100). In order to more easily allow confirmation of previous work and as a basis for further work, information has been added concerning the genetic background of the parents that had been used for crossing. Thus, in addition to the species involved, the cultivar-group (for C. pepo), market type (for C. maxima, C. moschata), and/or cultivar name are included in the description wherever possible. Genes affecting phenotypic/morphological traits are listed in Table 1. The data upon which are based identifications and concomitant assignment of gene symbols vary considerably in their content. No attempt is made here to assess the certainty of identifications, but gene symbols have been accepted or assigned only for cases in which at least some data are presented. The genes that are protein/isozyme variants are listed in Table 2. It can be seen from Tables 1 and 2 that a large number of genes, 65, have been identified for C. pepo L. For C. moschata Duchesne and C. maxima Duchesne, 21 and 19 genes have been identified, respectively, and for the interspecific cross of C. maxima × C. ecuadorensis Cutler & Whitaker, 29, of which 25 are isozyme variants. One or two genes have also been identified in four of the wild species (C. okeechobeensis Bailey, C. lundelliana Bailey, C. foetidissima HBK and C. ecuadorensis) and in several other interspecific crosses. Notably, no genes have been identified for the other two cultivated species, C. argyrosperma Huber and C. ficifolia Bouché. Some genes are listed as occurring in more than one species. This does not necessarily indicate that these genes reside at identical locations in the genome of different species. New additions to the list of Cucurbita genes include a number of omissions as well as a number of new genes published after the last update. Those that had been omitted are: Bn, pm-1, pm-2, and s-2, and Wmv ecu . Those that have been published since the last update are: Cmv, grl, l-1 BSt , l-1 iSt , L-2 w , m- zym mos , prv, qi, sl, wmv, zym mos , Zym-2, and Zym-3. In addition, there are many additions to the list of isozyme variants.

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Page 1: Gene List for Cucurbita species, 2004 - Cucurbit Breedingcuke.hort.ncsu.edu/cgc/cgcgenes/gene04squash.pdf · Gene List for Cucurbita species, 2004 Harry S. Paris A.R.O., Newe Ya’ar

Gene List for Cucurbita species, 2004 Harry S. Paris A.R.O., Newe Ya’ar Research Center, Ramat Yishay 30-095 (Israel) Rebecca Nelson Brown Oregon State University, Corvallis, OR 97331 (U.S.A.) A complete list of genes for Cucurbita species was last published 12 years ago (33). Since then, only updates have been published (72, 73). The genus Cucurbita L. contains 12 or 13 species (50). As far as is known, all have a complement of 20 pairs of chromosomes (2n = 40). This new gene list for Cucurbita contains much more detail concerning the sources of information, being modeled after the one for cucumber presented by Wehner and Staub (96) and its update by Xie and Wehner (100). In order to more easily allow confirmation of previous work and as a basis for further work, information has been added concerning the genetic background of the parents that had been used for crossing. Thus, in addition to the species involved, the cultivar-group (for C. pepo), market type (for C. maxima, C. moschata), and/or cultivar name are included in the description wherever possible. Genes affecting phenotypic/morphological traits are listed in Table 1. The data upon which are based identifications and concomitant assignment of gene symbols vary considerably in their content. No attempt is made here to assess the certainty of identifications, but gene symbols have been accepted or assigned only for cases in which at least some data are presented. The genes that are protein/isozyme variants are listed in Table 2. It can be seen from Tables 1 and 2 that a large number of genes, 65, have been identified for C. pepo L. For C. moschata Duchesne and C. maxima Duchesne, 21 and 19 genes have been identified, respectively, and for the interspecific cross of C. maxima × C. ecuadorensis Cutler & Whitaker, 29, of which 25 are isozyme variants. One or two genes have also been identified in four of the wild species (C. okeechobeensis Bailey, C. lundelliana Bailey, C. foetidissima HBK and C. ecuadorensis) and in several other interspecific crosses. Notably, no genes have been identified for the other two cultivated species, C. argyrosperma Huber and C. ficifolia Bouché. Some genes are listed as occurring in more than one species. This does not necessarily indicate that these genes reside at identical locations in the genome of different species. New additions to the list of Cucurbita genes include a number of omissions as well as a number of new genes published after the last update. Those that had been omitted are: Bn, pm-1, pm-2, and s-2, and Wmvecu. Those that have been published since the last update are: Cmv, grl, l-1BSt, l-1iSt, L-2w, m-zymmos, prv, qi, sl, wmv, zymmos, Zym-2, and Zym-3. In addition, there are many additions to the list of isozyme variants.

Page 2: Gene List for Cucurbita species, 2004 - Cucurbit Breedingcuke.hort.ncsu.edu/cgc/cgcgenes/gene04squash.pdf · Gene List for Cucurbita species, 2004 Harry S. Paris A.R.O., Newe Ya’ar

Symbols of genes that have been published in previous lists but have been modified for this list are Pm (to be used solely for powdery mildew resistance in C. lundelliana, with the separate designation Pm-0 for resistance in and derived from C. okeechobeensis), and Zym (with separate designations for different sources of resistance, viz. zymecu from C. ecuadorensis, Zym-0 from C. moschata ‘Nigerian Local’, and Zym-1 from C. moschata ‘Menina’, and zymmos from C. moschata ‘Soler’). Before choosing a gene name and symbol, researchers are urged to consult this Gene List as well as the rules of Gene Nomenclature for the Cucurbitaceae that appears near the end of this Cucurbit Genetics Cooperative Report in order to avoid confusion arising from duplication of gene names and symbols. Please contact us if you find omissions or errors in this Gene List. Several cases of genetic linkage have been reported: D – mo-2 (56) and M – Wt (C. pepo) (66) and Bi – Lo-2 (C. ecuadorensis × C. maxima) (30). Some of the isozyme variants observed by Weeden & Robinson (95) were also found to be linked to one another. RAPD markers have been categorized and organized into linkage groups and are not listed here but can be found in Brown and Myers (4) and Zraidi and Lelley (101). These two maps cannot be easily compared, as they were constructed using different mapping populations; RAPD markers are population-specific. Neither map gives complete coverage of the Cucurbita genome. Both maps contain morphological traits, either as single genes or as quantitative trait loci (QTLs). These traits are listed in Table 3 along with the most tightly linked RAPD markers. In many species, knowledge of the genome has moved beyond mapping markers linked to phenotypic traits to isolating and sequencing the genes that control the traits. Sequenced genes can be valuable to breeders and geneticists, as the differences in the gene sequences that result in the phenotypes of interest can be used as markers in marker-assisted selection. Unlike random markers, these gene-specific, allele-specific markers are completely linked to the genes of interest. Genes can be isolated through widespread sequencing of genomic or cDNA libraries, through map-based cloning, or by functional homology with sequenced genes from other species. In addition, genes which code for a known protein such as an enzyme can be isolated by working backwards from the protein. Many of the genes sequenced in Cucurbita at present have been sequenced this way. Map-based cloning is the most effective way to identify the DNA sequence of genes for phenotypic and morphological traits. This requires maps of much higher resolution than those presently available for Cucurbita. Most of the genes sequenced in Cucurbita have been isolated by researchers doing comparative studies of specific genes across plant families; usually only a single allele is available. Nonetheless, we have included a list of the sequenced genes in Table 4 as the sequences could be useful as a starting point for breeders interested in isolating the genes from lines of differing phenotype. In addition to the genes listed here, there exists a collection of partial sequences from mRNA for genes differentially expressed during seed development in C. pepo. These expressed sequence tags were identified in a study of the naked seed trait. The Gene Accession numbers for these sequences are CD726806 through CD726832.

Page 3: Gene List for Cucurbita species, 2004 - Cucurbit Breedingcuke.hort.ncsu.edu/cgc/cgcgenes/gene04squash.pdf · Gene List for Cucurbita species, 2004 Harry S. Paris A.R.O., Newe Ya’ar

Table 1. Phenotypic/Morphological Characteristics Gene Symbol

Preferred Synonym Character Species Reference(s)

a androecious. Found in ‘Greckie’; produces only male flowers, recessive to A.

pepo 39

B Bicolor. Precocious yellow fruit pigmentation; pleiotropic, affecting fruit and foliage, modified by Ep-1, Ep-2 and Ses-B. Originally from ‘Vaughn’s Pear Shaped’ ornamental gourd. B in C. moschata ‘Precocious PI 165561’ derived from C. pepo through backcrossing. Complementary to L-2 for intense orange, instead of light yellow, fruit-flesh color.

pepo, moschata 52, 68, 78, 85, 87

Bmax B-2 Bicolor. Precocious yellow fruit pigmentation, from subsp. andreana PI 165558

maxima 86, 89

Bi Bitter fruit. High cucurbitacin content in fruit. Bi from C. maxima subsp. andreana and C. ecuadorensis; bi from C. maxima subsp. maxima, including ‘Queensland Blue’. Linked to Lo-2. In C. pepo, Bi from wild Texan gourd; bi from zucchini squash.

maxima, maxima × ecuadorensis, pepo

11, 28, 30

bl blue fruit color. Incompletely recessive to Bl for green fruit color, in hubbard squash.

maxima 31

Bn* Butternut fruit shape, from ‘New Hampshire Butternut’, dominant to bn for crookneck fruit shape, as in ‘Canada Crookneck’.

moschata 48

Bu Bush habit. Short internodes; dominant to vine habit, bu, in young plant stage. In C. pepo, Bu in ‘Giant Yellow Straightneck’ and near-isogenic line of ‘Table Queen’, bu in ‘Table Queen’ acorn. In C. maxima, Bu from inbred line, bu from ‘Delicious’.

pepo, maxima 17, 29, 84

Cmv Cucumber mosaic virus resistance, from Nigerian Local. Dominant to cmv for susceptiblity, from ‘Waltham Butternut’.

moschata 3

cr cream corolla. Cream to nearly white petals, cr from C. okeechobeensis; Cr from C. moschata ‘Butternut’ incompletely dominant (yellow petals for Cr/cr, and orange for Cr/Cr)

moschata × okeechobeensis

75

cu cucurbitacin-B reduced; cu for reduced cucurbitacin-B content of cotyledons of ‘Early Golden Bush Scallop’; Cu for high cucurbitacin content of cotyledons of ‘Black Zucchini’.

pepo 83

D Dark stem. Series of three alleles observed in C. pepo: D for dark stem and dark intermediate-age

pepo, maxima 25, 42, 55, 56, 59, 67, 80

Page 4: Gene List for Cucurbita species, 2004 - Cucurbit Breedingcuke.hort.ncsu.edu/cgc/cgcgenes/gene04squash.pdf · Gene List for Cucurbita species, 2004 Harry S. Paris A.R.O., Newe Ya’ar

fruit, Ds for dark stem but fruit not affected, and d for light stem and fruit not affected, with dominance D > Ds > d. D from ‘Fordhook Zucchini’, Ds from ‘Early Prolific Straightneck’; d from ‘Vegetable Spaghetti’. Epistatic to genes l-1 and l-2 when either is homozygous recessive; linked to mo-2. In C. maxima, only the fruit was observed: D for dark intermediate-age fruit from the zapallito ‘La Germinadora’; d for light intermediate-age fruit from a variant zapallito breeding stock.

de determinate plant habit; stem lacking tendrils and terminating with female flowers. Recessive to De for indeterminate plant habit. De from ‘Jeju’ and ‘Sokuk’, de from inbred designated “Det”.

moschata 40

Di Disc fruit shape. From scallop squash, dominant to spherical or pyriform.

pepo 91, 97

Ep-1 Extender of pigmentation-1; modifier of B. Ep-1 incompletely dominant to ep-1 and additive with Ep-2. Ep-1 from ‘Small Sugar 7 × 7’ pumpkin; ep-1 from ‘Table King’ acorn.

pepo 90

Ep-2 Extender of pigmentation-2; modifier of B. Ep-2 incompletely dominant to ep-2 and additive with Ep-1. Ep-2 from ‘Table King’ acorn; ep-2 from ‘Small Sugar 7 × 7’ pumpkin.

pepo 90

Fr Fruit fly (Dacus cucurbitae) resistance. Fr from ‘Arka Suryamukhi’, dominant to fr for susceptibility.

maxima 49

fv fused vein. Fusion of primary leaf veins, subvital male gametophyte; found in hull-less-seeded pumpkin breeding line.

pepo 7, 8

G a, m Gynoecious sex expression; dominant to g for monoecious sex expression.

foetidissima 18, 23

Gb Green band on inner side of base of petal, from a scallop squash; dominant to gb, for no band, from a straightneck squash.

pepo 19

gc green corolla. Green, leaf-like petals, sterile; in unspecified F2 population.

pepo 92

gl glabrous, lacking trichomes maxima 37

Gr G Green rind. Dominant to buff skin of mature fruit. Gr from ‘Long Neapolitan’, gr from ‘Butternut’.

moschata 71

grl gray leaf. Recessive to green leaf. Recessive grl derived from cross of zapallito-type line of C. maxima and a butternut-type line of C. moschata. Dominant Grl from zapallito-type C. maxima.

maxima × moschata

41

Hi Hard rind inhibitor. Hi, for hard-rind inhibition, from C. maxima ‘Queensland Blue’; hi, for no hard-rind inhibition, from C. ecuadorensis.

maxima × ecuadorensis

30

Page 5: Gene List for Cucurbita species, 2004 - Cucurbit Breedingcuke.hort.ncsu.edu/cgc/cgcgenes/gene04squash.pdf · Gene List for Cucurbita species, 2004 Harry S. Paris A.R.O., Newe Ya’ar

Hr Hard rind. Hr for hard (lignified) rind in ornamental gourd, straightneck squash, and zucchini; hr for soft (non-lignified) rind in ‘Small Sugar’ pumpkin and ‘Sweet Potato’ (‘Delicata’). Complementary to Wt for Warty fruit.

pepo 44, 79

i intensifier of the cr gene for cream flowers. Cr/__ I/__ for intense orange or yellow flowers, Cr/__ i/i for light orange or yellow flowers, cr/cr I/__ for cream flowers, cr/cr i/i for white flowers. I from C. moschata ‘Butternut’, i from C. okeechobeensis.

moschata × okeechobeensis

75

I-mc Imc Inhibitor of mature fruit color; dominant to i-mc for no inhibition. I-mc in a scallop squash.

pepo 9

I-T Inhibitor of the T gene for trifluralin resistance. I-T from ‘La Primera’; i-t from ‘Ponca’ and ‘Waltham Butternut’.

moschata 1

l-1 c, St light fruit coloration-1. Light intensity of fruit coloration. Series of five alleles observed in C. pepo which, in complementary interaction with the dominant L-2 allele, give the following results: L-1 for uniformly intense/dark fruit coloration, from ‘Fordhook Zucchini’; l-1BSt for broad, contiguous intense/dark stripes, from ‘Cocozelle’; l-1St for narrow, broken intense/dark stripes, from ‘Caserta’; l-1iSt for irregular intense/dark stripes, from ‘Beirut’ vegetable marrow; l-1 for light coloration, from ‘Vegetable Spaghetti’, with dominance of L-1 > (l-1BSt > l-1St) ≥ l-1iSt > l-1. In C. maxima, L-1 from the zapallito ‘La Germinadora’; l-1 from a variant zapallito breeding stock.

pepo, maxima 25, 42, 57, 62, 63, 67, 76, 85

l-2 r light fruit coloration-2. Light intensity of fruit coloration. Series of three alleles observed in C. pepo, which, in complementary interaction with dominant alleles at the l-1 locus, give the following results: L-2 for intense/dark fruit coloration, with L-1 from ‘Fordhook Zucchini’ and intense/dark fruit stripes, with l-1BSt from ‘Cocozelle’; allele L-2w has delayed and weaker effect than L-2, from C. pepo subsp. fraterna; l-2 for light coloration, from ‘Vegetable Spaghetti’, with dominance of L-2 > L-2w > l-2. Dominant L-2 is also complementary with B for intense orange, instead of light yellow, fruit-flesh color and with recessive qi for intense exterior color of young fruit. In C. maxima, L-2 from the zapallito ‘La Germinadora’; l-2 from a variant zapallito breeding stock.

pepo, maxima 25, 42, 52, 58, 60, 67

lo-1 l lobed leaves-1; recessive to Lo-1 for non-lobed leaves

maxima 20

Lo-2 Lobed leaves-2. Lo-2 for lobed leaves in C. ecuadorensis dominant to lo-2 for unlobed leaves in

ecuadorensis × maxima

30

Page 6: Gene List for Cucurbita species, 2004 - Cucurbit Breedingcuke.hort.ncsu.edu/cgc/cgcgenes/gene04squash.pdf · Gene List for Cucurbita species, 2004 Harry S. Paris A.R.O., Newe Ya’ar

C. maxima. Linked to Bi.

lt leafy tendril. Tendrils with laminae; lt found in ornamental gourd.

pepo 77

ly light yellow corolla. Recessive to orange yellow; ly found in ornamental gourd.

pepo 77

M Mottled leaves. M for silver-gray areas in axils of leaf veins, dominant to m for absence of silver-gray. For C. maxima, M in ‘Zuni’ and m in ‘Buttercup’ and ‘Golden Hubbard’. For C. pepo, M in ‘Caserta’ and inbred of ‘Striato d’Italia’ cocozelle; m in ‘Early Prolific Straightneck’ and ‘Early Yellow Crookneck’. For C. moschata, M in ‘Hercules’ and ‘Golden Cushaw’, m in butternut type. Weakly linked to Wt.

pepo, maxima, moschata

13, 66, 76, 81

Mldg Mottled light and dark green immature fruit color; germplasm unspecified. Dominant to mldg for non-mottled.

moschata 5

mo-1 mature orange-1; complementary recessive gene for loss of green fruit color prior to maturity. Mo-1 from ‘Table Queen’ acorn; mo-1 from ‘Vegetable Spaghetti’.

pepo 56

mo-2 mature orange-2; complementary recessive gene for loss of green fruit color prior to maturity. Mo-2 from ‘Table Queen’ acorn; mo-2 from ‘Vegetable Spaghetti’. Linked to D.

pepo 56

ms-1 ms1 male sterile-1. Male flowers abort before anthesis, derived from a cross involving ‘Golden Hubbard’, recessive to Ms-1 for male fertile.

maxima 82

ms-2 ms2 male sterile-2. Male flowers abort, sterility expressed as androecium shrivelling and turning brown; ms-2 from ‘Eskandarany’ (PI 228241).

pepo 22

ms-3 ms-2 male sterile-3. maxima 37

m-zymmos* modifier of dominance of zucchini yellow mosaic virus resistance; confers resistance to otherwise susceptible Zymmos/zymmos heterozygotes. M-zymmos in ‘Soler’, m-zymmos in ‘Waltham Butternut’ and ‘Nigerian Local’.

moschata 51

n h naked seeds. Lacking a lignified seed coat, n from oil-seed pumpkin.

pepo, moschata 27, 80, 87, 99, 102

pl plain light fruit color, pl from ‘Beirut’ vegetable marrow and ‘Fordhook Zucchini’; Pl in ‘Vegetable Spaghetti’.

pepo 53

Pm, Pm-0*

Powdery mildew resistance. Resistance to Podosphaera xanthii; Pm from C. lundelliana; Pm-0 from C. okeechobeensis and in C. pepo

lundelliana, okeechobeensis, pepo

10, 12, 35, 70

Page 7: Gene List for Cucurbita species, 2004 - Cucurbit Breedingcuke.hort.ncsu.edu/cgc/cgcgenes/gene04squash.pdf · Gene List for Cucurbita species, 2004 Harry S. Paris A.R.O., Newe Ya’ar

pm-1 powdery mildew resistance in C. moschata. Series of three alleles: pm-1P for susceptibility from ‘Ponca’ dominant to pm-1L for resistance from ‘La Primera’, which is dominant to pm-1W for susceptibility in ‘Waltham Butternut’.

moschata 2

pm-2 powdery mildew resistance in C. moschata ‘Seminole’, recessive to Pm-2 for susceptibility

moschata 2

prv papaya ringspot virus resistance, in Nigerian Local, recessive to Prv for susceptibility, in ‘Waltham Butternut’.

moschata 3

qi quiescent intense. Recessive to Qi for not intense and complementary to L-2 for intense young fruit color; little or no effect on mature fruit. Qi from ‘Vegetable Spaghetti’; qi from ‘Jack O’Lantern’ pumpkin and ‘Verte non-coureuse d’Italie’ cocozelle.

pepo 58, 61

Rd Red skin. Red external fruit color; dominant to green, white, yellow and gray. Rd from ‘Turk’s Cap’; rd from ‘Warted Hubbard’.

maxima 43

ro rosette leaf. Lower lobes of leaves slightly spiraled, ro derived from an ornamental gourd.

pepo 44

s-1 s sterile. Male flowers small, without pollen; female flower sterile. Derived from crossing ‘Greengold’ with ‘Banana’.

maxima 32

s-2 sterile. Male flowers small, without pollen and female flower sterile; mutant in powdery mildew resistant, straightneck squash breeding line.

pepo 6

Ses-B Selective suppression of gene B. Suppression in foliage of precocious yellowing conferred by B. Ses-B in straightneck breeding line dominant to ses-B in ‘Jersey Golden Acorn’.

pepo 88

sl silverleaf resistance. Recessive to Sl for susceptibility. Sl from ‘Soler’; sl from PI 162889 and butternut types.

moschata 26

Slc Squash leaf curl virus resistance; derived from C. moschata.

pepo 46

sp spaghetti flesh, breaking into strands after cooking pepo 45

T Trifluralin resistance. Dominant to susceptibility to the herbicide; modified by I-T. T in ‘La Primera’; t in ‘Ponca’ and ‘Waltham Butternut’.

moschata 1

uml umbrella-like; leaves shaped like partially opened umbrella. Recessive uml derived from a cross of C. maxima ‘Warzywna’ and a C. pepo inbred; dominant Uml from ‘Warzywna’.

maxima × pepo 69

v virescent. Yellow-green young leaves, v found in ‘Golden Delicious’.

maxima 21

Page 8: Gene List for Cucurbita species, 2004 - Cucurbit Breedingcuke.hort.ncsu.edu/cgc/cgcgenes/gene04squash.pdf · Gene List for Cucurbita species, 2004 Harry S. Paris A.R.O., Newe Ya’ar

W Weak fruit coloration. Dominant to w for intense-pigmented mature fruit; W from scallop squash. Complementary to Wf for white external fruit color.

pepo 54, 91, 97

wc white corolla. Derived from ‘Ispanskaya’ × ‘Emerald’. Recessive to Wc for normal orange-yellow corolla

maxima 38

Wf White flesh. Dominant to wf for colored flesh. Wf in a scallop squash, wf in a straightneck squash. Complementary to W for white external fruit color.

pepo 19, 54, 91

Wmv Watermelon mosaic virus resistance. From “Menina” and “Nigerian Local”, dominant to wmv for susceptibility in ‘Musquée de Provence’ and ‘Waltham Butternut’. May be linked with or identical to Zym-1.

moschata 3, 24

Wmvecu* Watermelon mosaic virus resistance. From C. ecuadorensis, in a cross with an unspecified C. maxima.

maxima × ecuadorensis

95

Wt Warty fruit. Dominant to non-warted, wt, and complementary to Hr, with fruit wartiness being expressed only in the presence of the dominant Hr allele. Wt in straightneck, crookneck, and ‘Delicata’; wt in zucchini, cocozelle, and ‘Small Sugar’ pumpkin. Weakly linked to M.

pepo 66, 79, 91

wyc white-yellow corolla; isolated in ‘Riesen-Melonen’. Recessive to Wyc for normal orange-yellow corolla.

maxima 38

Y Yellow fruit color. Y for yellow fruit color of intermediate-age fruits, from straightneck and crookneck squash, dominant to y for green intermediate-age fruit color, from vegetable marrow, ornamental gourd, and cocozelle.

pepo 66, 76, 84, 85, 91

yg yellow-green leaves and stems maxima 37

Ygp Yellow-green placenta. Dominant to yellow placental color. Ygp in a scallop squash, ygp in a straightneck squash.

pepo 19

ys yellow seedling. Lacking chlorophyll; lethal pepo 44

zymecu zucchini yellow mosaic virus resistance, recessive to susceptibility; zymecu from C. ecuadorensis, Zymecu from C. maxima ‘Buttercup’.

ecuadorensis 74

zymmos* zucchini yellow mosaic virus resistance, recessive to susceptibility; zymmos from ‘Soler’, Zymmos from ‘Waltham Butternut’.

moschata 51

Zym-0* Zucchini yellow mosaic virus resistance. Zym-0 from C. moschata ‘Nigerian Local’ dominant to zym-0 for susceptibility from ‘Waltham Butternut’. Perhaps one of two separate genes for resistance in ‘Nigerian Local’.

moschata 3, 47, 51

Page 9: Gene List for Cucurbita species, 2004 - Cucurbit Breedingcuke.hort.ncsu.edu/cgc/cgcgenes/gene04squash.pdf · Gene List for Cucurbita species, 2004 Harry S. Paris A.R.O., Newe Ya’ar

Zym-1 Zucchini yellow mosaic virus resistance. Zym-1 from C. moschata ‘Menina’ dominant to zym-1 for susceptibility from C. moschata ‘Waltham Butternut’. Zym-1 transferred via backcrossing to C. pepo ‘True French’ zucchini, in which it confers resistance through complementary interaction with Zym-2 and Zym-3. Zym-1 is either linked with Wmv or also confers resistance to watermelon mosaic virus.

moschata, pepo 24, 51, 64, 65

Zym-2 Zucchini yellow mosaic virus resistance-2. Dominant to susceptibility and complementary to Zym-1. Zym-2 from C. moschata ‘Menina’. Zym-2 in C. pepo derived from C. moschata, in near-isogenic resistant line of ‘True French’ zucchini; zym-2 from C. pepo ‘True French’.

moschata, pepo 64

Zym-3 Zucchini yellow mosaic virus resistance-3. Dominant to susceptibility and complementary to Zym-1. Zym-3 from C. moschata ‘Menina’. Zym-3 in C. pepo derived from C. moschata, in near-isogenic resistant line of ‘True French’ zucchini; zym-3 from C. pepo ‘True French’.

moschata, pepo 64

*Proposed new gene symbol. Table 2. Isozyme Variants

Gene Symbol Preferred Synonym No. alleles

observed Character Species Reference(s)

Aat-1 Aat 8 Aspartate aminotransferase-1. Variant among accessions.

pepo 16, 34

Aat-3 2 Aspartate aminotransferase-3. Variant among wild populations.

pepo 16

Aat-4 3 Aspartate aminotransferase-4. Variant among wild populations.

pepo 16

Aat-mb 2 Aspartate aminotransferase – microbody

maxima × ecuadorensis

95

Aat-m1 2 Aspartate aminotransferase mitochondria-1

maxima × ecuadorensis

95

Aat-m2 2 Aspartate aminotransferase mitochondria-2

maxima × ecuadorensis

95

Aat-p2 2 Aspartate aminotransferase plastid-2 maxima × ecuadorensis

95

Acp-1 2 Acid phosphatase-1 maxima × 95

Page 10: Gene List for Cucurbita species, 2004 - Cucurbit Breedingcuke.hort.ncsu.edu/cgc/cgcgenes/gene04squash.pdf · Gene List for Cucurbita species, 2004 Harry S. Paris A.R.O., Newe Ya’ar

ecuadorensis Acp-2 2 Acid phosphatase-2 maxima ×

ecuadorensis 95

Aldo-p 2 Aldolase – plastid maxima × ecuadorensis

94

Est-1 Est 2 Esterase maxima × ecuadorensis

93, 95

Gal-1 2 β-galactosidase-1 maxima × ecuadorensis

95

Gal-2 2 β-galactosidase-2 maxima × ecuadorensis

95

G2d-1 3 Glycerate dehydrogenase-1. Variant among wild populations.

pepo 16

G2d-2 2 Glycerate dehydrogenase-2. Variant among wild populations.

pepo 16

Got-1 5 Glutamine oxaloacetate-1. Variant among accessions, wild populations, and among Cucurbita species.

pepo 14, 15, 36, 98

Got-2 3 Glutamine oxaloacetate-2. Variant among species.

maxima × ecuadorensis

98

Gpi 2 Glucosephosphate isomerase. Variant among accessions.

pepo 34

Gpi-3 2 Glucosephosphate isomerase-3. Variant among wild populations.

pepo 16

Gpi-c1 2 Glucosephosphate isomerase cytosolic-1

maxima × ecuadorensis

95

Gpi-c2 2 Glucosephosphate isomerase cytosolic-2

maxima × ecuadorensis

95

Idh-1 4 Isocitrate dehydrogenase-1. Variant among accessions, wild populations, and Cucurbita species.

pepo 14, 15, 16, 36, 98

Idh-2 2 Isocitrate dehydrogenase-2. Variant among accessions, wild populations, and Cucurbita species.

pepo 14, 15, 16, 36, 98

Idh-3 2 Isocitrate dehydrogenase-3. Variant among accessions and populations.

pepo 14, 15, 16, 36

Lap-1 Lap 4 Leucine aminopeptidase. Variant among C. pepo accessions.

maxima × ecuadorensis; pepo

16, 34 , 93, 95

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Mdh-1 Mdh 7 Malate dehydrogenase. Variant among accessions.

pepo 34

Mdh-2 3 Malate dehydrogenase-2. Variant among accessions, wild populations, and Cucurbita species.

pepo 14, 15, 16, 36, 98

Mdh-3 3 Malate dehydrogenase-3. Variant among accessions, wild populations, and Cucurbita species.

pepo 14, 15, 16, 36, 98

Mdh-m1 2 Malate dehydrogenase mitochondria-1

maxima × ecuadorensis

95

Mdh-m2 2 Malate dehydrogenase mitochondria-2

maxima × ecuadorensis

95

Mdh-c2 2 Malate dehydrogenase cytosolic-2 maxima × ecuadorensis

95

Per-1 2 Peroxidase-1 maxima × ecuadorensis

95

Per-2 3 Peroxidase-2. Variant among accessions and wild populations.

pepo 14, 15, 36

Per-3 2 Peroxidase-3 maxima × ecuadorensis

95

Pgi-1 2 Phosphoglucase isomerase-1 pepo 14

Pgi-2 2 Phosphoglucase isomerase-2. Variant among Cucurbita species.

pepo 14, 36, 98

Pgi-3 4 Phosphoglucase isomerase-3. Variant among accessions, wild populations, and Cucurbita species.

pepo 14, 15, 36, 98

Pgm-1 Pgm 2 Phosphoglucomutase. Variant among accessions.

pepo 34

Pgm-2 4 Phosphoglucomutase-2. Variant among accessions, wild populations, and Cucurbita species.

pepo 14, 15, 36, 98

Pgm-5 2 Phosphoglucomutase-5. Variant among wild populations.

pepo 16

Pgm-6 2 Phosphoglucomutase-6. Variant among wild populations.

pepo 16

Pgm-c2 2 Phosphoglucomutase cytosolic-2 maxima × ecuadorensis

95

Pgm-p 2 Phosphoglucomutase plastid maxima × ecuadorensis

95

Skd-1 6 Shikimate dehydrogenase. Variant pepo 16

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among wild populations.

Skdh 5 Shikimate dehydrogenase. Variant among C. pepo accessions.

maxima × ecuadorensis; pepo

34, 95

Sod-1 2 Superoxide dismutase-1 maxima × ecuadorensis

95

Tpi-c2 2 Triosephosphatase isomerase cytosolic-2

maxima × ecuadorensis

95

Tpi-p2 2 Triosephosphatase isomerase plastid-2

maxima × ecuadorensis

95

Table 3. Mapped Phenotypic/Morphological Characteristics Trait Symbol Linked Marker(s) Recombination

Distance (cM) Reference(s)

Seed Coat n AK11_340 4.4 101 Fruit Length (QTL) AE07_165, AC10_490, AJ20_420, P13_750,

J01_600, AO20_1200, T08_460, AB08_540, AE09_1600

101

Fruit Width (QTL) AE07_165, AJ20_420, AM10_950, AG08_440

101

Fruit Length/width Ratio

(QTL) AE07_165, AC10_490, AJ20_420, P13_750, J01_600

101

No. of Fruit Chambers

(QTL) P13_950, AE08_470 101

Precocious yellow fruit

B I10_1700 27.1 4

Leaf Indentation

(QTL) F10_400, K11_950, G2_400 4

Leaf Mottle M H14_600 U489_1200

13.0 16.3

4

Mature Fruit Color

[none given]

G17_700 9.7 4

Fruit Shape (QTL) F8_1050, B8_900, H19_500 4

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Literature Cited in text and Tables 1, 2, and 3 1. Adeniji, A.A. and D.P. Coyne. 1981. Inheritance of resistance to trifluralin toxicity in Cucurbita moschata Poir.

HortScience 16: 774–775. 2. Adeniji, A.A. and D.P. Coyne. 1983. Genetics and nature of resistance to powdery mildew in crosses of butternut with

calabaza squash and ‘Seminole Pumpkin’. J. Amer. Soc. Hort. Sci. 108: 360–368. 3. Brown, R.N., A. Bolanos-Herrera, J.R. Myers, and M.M. Jahn. 2003. Inheritance of resistance to four cucurbit viruses

in Cucurbita moschata. Euphytica 129: 253–258. 4. Brown, R.N. and J.R. Myers. 2002. A genetic map of squash (Cucurbita sp.) with randomly amplified polymorphic

DNA markers and morphological markers. J. Amer. Soc. Hort. Sci. 127: 568–575. 5. Cardosa, A.I.I., P.T. Della Vecchia, and N. Silva. 1993. Inheritance of immature fruit color in C. moschata. Cucurbit

Genet. Coop. Rep. 16: 68–69. 6. Carle, R.B. 1997. Bisex sterility governed by a single recessive gene in Cucurbita pepo. Cucurbit Genet. Coop. Rep.

20: 46–47. 7. Carle, R.B. and J.B. Loy. 1996. Genetic analysis of the fused vein trait in Cucurbita pepo L. J. Amer. Soc. Hort. Sci.

121: 13–17. 8. Carle, R.B. and J.B. Loy. 1996. Fused vein trait in Cucurbita pepo L. associated with subvitality of the male

gametophyte. J. Amer. Soc. Hort. Sci. 121: 18–22. 9. Clayberg, C.D. 1992. Reinterpretation of fruit color inheritance in Cucurbita pepo L. Cucurbit Genet. Coop. Rep. 15:

90–92. 10. Cohen, R., A. Hanan, and H.S. Paris. 2003. Single-gene resistance to powdery mildew in zucchini squash (Cucurbita

pepo). Euphytica 130: 433–441. 11. Contardi, H.G. 1939. Estudios geneticos en Cucurbita y consideraciones agronomicas. Physis 18: 331–347. 12. Contin, M. 1978. Interspecific transfer of powdery mildew resistance in the genus Cucurbita. Ph.D. Thesis, Cornell

Univ., Ithaca, New York. 13. Coyne, D.P. 1970. Inheritance of mottle-leaf in Cucurbita moschata Poir. HortScience 5: 226–227. 14. Decker, D.S. 1985. Numerical analysis of variation in Cucurbita pepo. Econ. Bot. 39: 300–309. 15. Decker, D.S. and H.D. Wilson. 1987. Allozyme variation in the Cucurbita pepo complex: C. pepo var. ovifera vs. C.

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18: 249–255. 26. Gonzalez-Roman, M. and L. Wessel-Beaver. 2002. Resistance to silverleaf disorder is controlled by a single recessive

gene in Cucurbita moschata Duchesne. Cucurbit Genet. Coop. Rep. 25: 49–50. 27. Grebenščikov, I. 1954. Zur Vererbung der Dünnschaligkeit bei Cucurbita pepo L. Züchter 24: 162–166. 28. Grebenščikov, I. 1955. Notulae cucurbitologicae II. Über Cucurbita texana A. Gr. und ihre Kreuzung mit einer

hochgezüchteten C. pepo-Form. Kulturpflanze 3: 50–59. 29. Grebenščikov, I. 1958. Notulae cucurbitologicae III. Kulturpflanze 6: 38–60.

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30. Herrington, M.E. and J.P. Brown. 1988. Inheritance of leaf and fruit characteristics in Cucurbita maxima Duch. cv. Queensland Blue × C. ecuadorensis Cutler and Whitaker. Queensl. J. Agr. Anim. Sci. 45: 45–48.

31. Hutchins, A.E. 1935. The interaction of blue and green color factors in hubbard squash. Proc. Amer. Soc. Hort. Sci. 33: 514.

32. Hutchins, A.E. 1944. A male and female sterile variant in squash, Cucurbita maxima Duch. Proc. Amer. Soc. Hort. Sci. 44: 494–496.

33. Hutton, M.G. and R.W. Robinson. 1992. Gene list for Cucurbita spp. Cucurbit Genet. Coop. Rep. 15: 102–109. 34. Ignart, F. and N.F. Weeden. 1984. Allozyme variation in cultivars of Cucurbita pepo L. Euphytica 33: 779–785. 35. Jahn, M., H.M. Munger, and J.D. McCreight. 2002. Breeding cucurbit crops for powdery mildew resistance. In: R.R.

Bélanger, W.R. Bushnell, A.J. Dik, and T.L.W. Carver, Eds., The powdery mildews, a comprehensive treatise, pp. 239–248. American Phytopathological Society, St. Paul, MN.

36. Kirkpatrick, K.J., D.S. Decker, and H.D. Wilson. 1985. Allozyme differentiation in the Cucurbita pepo complex: C. pepo var. medullosa vs. C. texana. Econ. Bot. 39: 289–299.

37. Korzeniewska, A. 1992. New genes in Cucurbita maxima Duch. In: R.W. Doruchowski, E. Kozik, and K. Niemirowicz-Szczytt, eds. Proc. Cucurbitaceae ’92: the 5th Eucarpia Meeting on Cucurbit Genetics & Breeding, pp. 75–78.

38. Korzeniewska, A. 1996. Two independent loci for white and white-yellow corolla in Cucurbita maxima Duch. In: M.L. Gomez-Guillamon, C. Soria, J. Cuartero, J. Tores, and R. Fernandez-Munoz, eds. Proc. Cucurbitaceae Towards 2000: The 6th Eucarpia Meeting on Cucurbit Genetics & Breeding. Graficas Axarquia, Velez-Malaga, Spain, pp. 78–81.

39. Kubicki, B. 1970. Androecious strains of Cucurbita pepo L. Genet. Polon. 11: 45–51. 40. Kwack, S.N. 1995. Inheritance of determinate growth habit in Cucurbita moschata Poir. J. Kor. Soc. Hort. Sci. 36:

780–784. 41. Lopez-Anido, F., E. Cointry, I. Firpo, S.M. Garcia, and S. Gattuso. 2002. Inheritance of gray leaf color in a material

derived from a Cucurbita maxima Duch. × C. moschata Duch. hybrid. Cucurbit Genet. Coop. Rep. 25: 46–48. 42. Lopez-Anido, F., V. Cravero, P. Asprelli, E. Cointry, I. Firpo, and S.M. Garcia. 2003. Inheritance of immature fruit

color in Cucurbita maxima var. Zapallito (Carrière) Millan. Cucurbit Genet. Coop. Rep. 26: 48–50. 43. Lotsy, J.P. 1920. Cucurbita Strijdvragen. II. Eigen Onderzoekingen. Genetica 2: 1–21. 44. Mains, E.B. 1950. Inheritance in Cucurbita pepo. Papers Mich. Acad. Sci. Arts Letters 36: 27–30. 45. Mazurek, Z. and K. Niemirowicz-Szczytt. 1992. Inheritance of spaghetti traits in Cucurbita pepo. In: R.W.

Doruchowski, E. Kozik, and K. Niemirowicz-Szczytt, eds. Proc. Cucurbitaceae ’92: the 5th Eucarpia Meeting on Cucurbit Genetics & Breeding, pp. 70–74.

46. Montes-Garcia, C.E., S. Garza-Ortega, and J.K. Brown. 1998. Inheritance of the resistance to squash leaf curl virus in Cucurbita pepo L. In: J.D. McCreight, ed. Cucurbitaceae ’98: Evaluation and Enhancement of Cucurbit Germplasm. A.S.H.S., Alexandria, Virginia, pp. 328–330.

47. Munger, H.M. and R. Provvidenti. 1987. Inheritance of resistance to zucchini yellow mosaic virus in Cucurbita moschata. Cucurbit Genet. Coop. Rep. 10: 80–81.

48. Mutschler, M.A. and O.H. Pearson. 1987. The origin, inheritance, and instability of butternut squash (Cucurbita moschata Duchesne). HortScience 22: 535–539.

49. Nath, P., O.P. Dutta, S. Velayudhan, and K.R.M. Swamy. 1976. Inheritance of resistance to fruit fly in pumpkin. Sabrao J. 8: 117–119.

50. Nee, M. 1990. The domestication of Cucurbita (Cucurbitaceae). Econ. Bot. 44(3, Suppl.): 56–68. 51. Pachner, M. and T. Lelley. 2004. Different genes for resistance to zucchini yellow mosaic virus (ZYMV) in Cucurbita

moschata. In: A. Lebeda and H.S. Paris (eds.), Proceedings of Cucurbitaceae 2004, pp. 237–243. Palacký Univ., Olomouc, Czech Republic.

52. Paris, H.S. 1988. Complementary genes for orange fruit flesh color in Cucurbita pepo. HortScience 23: 601–603. 53. Paris, H.S. 1992. A recessive, hypostatic gene for plain light fruit coloration in Cucurbita pepo. Euphytica 60: 15–20. 54. Paris, H.S. 1995. The dominant Wf (white flesh) allele is necessary for expression of “white” mature fruit color in

Cucurbita pepo. In: G. Lester and J. Dunlap (Eds.), Cucurbitaceae ’94, pp. 219–220. Gateway, Edinburg, TX U.S.A. 55. Paris, H.S. 1996. Multiple allelism at the D locus in squash. J. Hered. 87: 391–395. 56. Paris, H.S. 1997. Genes for developmental fruit coloration of acorn squash. J. Hered. 88: 52–56. 57. Paris, H.S. 2000. Gene for broad, contiguous dark stripes in cocozelle squash. Euphytica 115: 191–196. 58. Paris, H.S. 2000. Quiescent intense (qi): a gene that affects young but not mature fruit color intensity in Cucurbita

pepo. J. Hered. 91: 333–339.

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59. Paris, H.S. 2000. Segregation distortion in Cucurbita pepo. In: N. Katzir and H.S. Paris (Eds.), Proceedings of Cucurbitaceae 2000. Acta Hort. 510: 199–202.

60. Paris, H.S. 2002. Multiple allelism at a major locus affecting fruit coloration in Cucurbita pepo. Euphytica 125: 149–153.

61. Paris, H.S. 2002. No segregation distortion in intersubspecific crosses in Cucurbita pepo. Cucurbit Genet. Coop. Rep. 25: 43–45.

62. Paris, H.S. 2003. Genetic control of irregular striping, a new phenotype in Cucurbita pepo. Euphytica 129: 119–126. 63. Paris, H.S. and Y. Burger. 1989. Complementary genes for fruit striping in summer squash. J. Hered. 80: 490–493. 64. Paris, H.S. and S. Cohen. 2000. Oligogenic inheritance for resistance to zucchini yellow mosaic virus in Cucurbita

pepo. Ann. Appl. Biol. 136: 209–214. 65. Paris, H.S., S. Cohen, Y. Burger, and R. Yoseph. 1988. Single gene resistance to zucchini yellow mosaic virus in

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H.S. Paris (Eds.), Proceedings of Cucurbitaceae 2004, pp. 389–394.. Palacký Univ., Olomouc, Czech Republic. 67. Paris, H.S. and H. Nerson. 1986. Genes for intense pigmentation of squash. J. Hered. 77: 403–409. 68. Paris, H.S., H. Nerson, and Y. Burger. 1985. Precocious PI 165561 and Precocious PI 165561R pumpkin breeding

lines. HortScience 20: 778–779. 69. Rakoczy-Trojanowska, M. and S. Malepszy. 1999. Inheritance of umbrella-like leaf shape in materials derived form

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HortScience 9: 464. 78. Schaffer, A.A. and C.D. Boyer. 1984. The influence of gene B on fruit development in Cucurbita pepo. J. Amer. Soc.

Hort. Sci. 106: 432–437. 79. Schaffer, A.A., C.D. Boyer, and H.S. Paris. 1986. Inheritance of rind lignification and warts in Cucurbita pepo L. and

a role for phenylalanine ammonia lyase in their control. Z. Pflanzenzüchtg. 96: 147–153. 80. Schöniger, G. 1952. Vorläufige Mitteilung über das Verhalten der Testa- und Farbgene bei verschiedenen Kreuzungen

innerhalb der Kürbisart Cucurbita pepo L. Züchter 22: 316–337. 81. Scott, D.H. and M.E. Riner. 1946. A mottled leaf character in winter squash. J. Hered. 37: 27–28. 82. Scott, D.H. and M.E. Riner. 1946. Inheritance of male sterility in winter squash. Proc. Amer. Soc. Hort. Sci. 47: 375–

377. 83. Sharma, G.C. and C.V. Hall. 1971. Cucurbitacin B and total sugar inheritance in Cucurbita pepo related to spotted

cucumber beetle feeding. J. Amer. Soc. Hort. Sci. 96: 750–754. 84. Shifriss, O. 1947. Developmental reversal of dominance in Cucurbita pepo. Proc. Amer. Soc. Hort. Sci. 50: 330–346. 85. Shifriss, O. 1955. Genetics and origin of the bicolor gourds. J. Hered. 46: 213–222. 86. Shifriss, O. 1966. Behavior of gene B in Cucurbita. Veg. Improv. Newsl. 8: 7–8. 87. Shifriss, O. 1981. Origin, expression, and significance of gene B in Cucurbita pepo L. J. Amer. Soc. Hort. Sci. 106:

220–232. 88. Shifriss, O. 1982. Identification of a selective suppressor gene in Cucurbita pepo L. HortScience 17: 637–638. 89. Shifriss, O. 1989. Relationship between the B genes of two Cucurbita species, II. Cucurbit Genet. Coop. Rep. 12: 75–

78. 90. Shifriss, O. and H.S. Paris. 1981. Identification of modifier genes affecting the extent of precocious fruit pigmentation

in Cucurbita pepo L. J. Amer. Soc. Hort. Sci. 106: 653–660. 91. Sinnott, E.W. and G.B. Durham. 1922. Inheritance in the summer squash. J. Hered. 13: 177–186. 92. Superak, T.H. 1987. A green corolla mutant in Cucurbita pepo. Cucurbit Genet. Coop. Rep. 10: 103.

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93. Wall, J.R. and T.W. Whitaker. 1971. Genetic control of leucine aminopeptidase and esterase isozymes in the interspecific cross Cucurbita ecuadorensis × C. maxima. Biochem. Genet. 5: 223–229.

94. Weeden, N.F., R.W. Robinson, and F. Ignart. 1984. Linkage between an isozyme locus and one of the genes controlling resistance to watermelon mosaic virus 2 in Cucurbita ecuadorensis. Cucurbit Genet. Coop. Rep. 7: 86–87.

95. Weeden, N.F. and R.W. Robinson. 1986. Allozyme segregation ratios in the interspecific cross Cucurbita maxima × C. ecuadorensis suggest that hybrid breakdown is not caused by minor alterations in chromosome structure. Genetics 114: 593–609.

96. Wehner, T.C. and J.E. Staub. 1997. 1997 gene list for cucumber. Cucurbit Genet. Coop. Rep. 20: 66–88. 97. Whitaker, T.W. 1932. Fertile gourd-pumpkin hybrids. J. Hered. 23: 427–430. 98. Wilson, H.D. 1989. Discordant patterns of allozyme and morphological variation in Mexican Cucurbita. Syst. Bot.

14: 612–623. 99. Xianglin, Z. 1987. A study on the breeding of naked kernel pumpkin and its genetic behavior. Acta Hort. Sin. 14:

115–118 (Chinese, with English summary). 100. Xie, J. and T.C. Wehner. 2001. Gene list 2001 for cucumber. Cucurbit Genet. Coop. Rep. 24: 110–136. 101. Zraidi, A. and T. Lelley. 2004. Genetic map for pumpkin Cucurbita pepo using random amplified polymorphic DNA

markers. In: A. Lebeda and H.S. Paris (Eds.), Proceedings of Cucurbitaceae 2004, pp. 507–514. Palacký Univ., Olomouc, Czech Republic.

102. Zraidi, A., M. Pachner, T. Lelley, and R. Obermayer. 2003. On the genetics and histology of the hull-less character of Styrian oil-pumpkin (Cucurbita pepo L.). Cucurbit Genet. Coop. Rep. 26: 57–61.

Table 4. Genes with known DNA sequence Gene Symbol*

Gene Accession

(Putative) Function Source Ref.

AIG-2 AY666083 aspartic protease inhibitor C. maxima **

PRB1 AY326308 phloem RNA-binding protein C. maxima ‘Big Max’ **

GAIP AY32630, AY326307

gibberellic acid insensitive phloem protein (two very similar genes)

C. maxima ‘Big Max’ **

FAD2 AY525163 omega-6 fatty acid desaturase C. pepo zucchini **

NIP1 AJ544830 Nod26-like protein C. pepo zucchini 23

PP2 AY312402 phloem protein 2 lectin (includes promoter region)

C. moschata crookneck **

PP2 AF150627 phloem protein 2 lectin C. moschata crookneck **

PP2 Z22647 phloem protein 2 lectin C. pepo ‘Autumn Gold’ 42

PP2 Z17331 phloem protein 2 lectin C. maxima ‘Big Max’ 2

PP2 L31550, L31551, L31552

phloem protein 2 (three alleles) C. maxima **

GA2OX, GA20OX, GA3OX

AJ315663, AJ302041, AJ308480, AJ302040

gibberellin oxidases (two sequences for GA2OX)

C. maxima ‘Riesenmelone’ **

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U61385 gibberellin 20-oxidase C. maxima ‘Riesenmelone’ 25

U63650 gibberellin 2 beta,3 beta hydroxylase C. maxima ‘Riesenmelone’ 26

AJ006453 gibberellin 3 beta hydroxylase C. maxima ‘Riesenmelone’ **

U61386 gibberellin dioxygenase C. maxima ‘Riesenmelone’ 24

Moschatin 1 through 5

AF462349, AF504011, AY25646, AY27921, AY279217

ribosome-inactivating protein C. moschata crookneck **

CPS1 AB109763 copalyl diphosphate synthase; gibberellin biosynthesis

C. maxima **

CPS AF049905, AF049906

copalyl diphosphate synthase; gibberellin biosynthesis (2 genes)

C. maxima 37

Hsc70 AF527794, AF527795, AF527796

cell-autonomous heat shock protein; chaperonin 70 (multiple sequences)

C. maxima 1

AB061204 thioredoxin h C. maxima **

Puga, Pugb, Pugc

AB055116, AB055117, AB055118

glutathione S-transferase C. maxima **

CYP88A AF212990, AF212991

cytochrome P450; ent-kaurenoic acid oxidase (multiple alleles)

C. maxima ‘Queensland Blue’ 13

PP2 AF520583 phloem protein 2 C. digitata PI 240879 **

PP2 AF520582 phloem lectin C. argyrosperma subsp. sororia **

L32700, L32701

phloem lectin C. argyrosperma 2

X56948 malate synthase Cucurbita sp.*** ‘Kurokawa Amakuri Nankin’

29

pMCPN60 X70867, X70868

chaperonin 60 ‘Kurokawa Amakuri Nankin’ 40

PCPK AY07280, AY072802

phloem calmodulin-like protein kinases C. maxima ‘Big Max’ 47

X55779 ascorbate oxidase C. maxima ‘Ebisu Nankin’ 7

AAO D55677 ascorbate oxidase C. maxima 21

chitP1 AB015655 chitinase C. maxima ‘Ebisu Nankin’ **

PLC AF082284 chitinase C. moschata crookneck 20

Page 18: Gene List for Cucurbita species, 2004 - Cucurbit Breedingcuke.hort.ncsu.edu/cgc/cgcgenes/gene04squash.pdf · Gene List for Cucurbita species, 2004 Harry S. Paris A.R.O., Newe Ya’ar

PV72 AB006809 vacuolar sorting receptor ‘Kurokawa Amakuri Nankin’ 36

D88420 stromal ascorbate peroxidase ‘Kurokawa Amakuri Nankin’ 28

D78256 isocitrate lyase ‘Kurokawa Amakuri Nankin’ 27

D70895 3-ketoacyl-CoA thiolase ‘Kurokawa Amakuri Nankin’ 19

D83656 thylakoid ascorbate peroxidase ‘Kurokawa Amakuri Nankin’ 45

D49433 hydroxypyruvate reductase ‘Kurokawa Amakuri Nankin’ 12

MP28 D45078 membrane protein ‘Kurokawa Amakuri Nankin’ 16

D38132 glyoxysomal citrate synthase ‘Kurokawa Amakuri Nankin’ 18

D29629 aconitase ‘Kurokawa Amakuri Nankin’ 10

D16560 prepro2S albumin ‘Kurokawa Amakuri Nankin’ 8

D14044 glycolate oxidase ‘Kurokawa Amakuri Nankin’ 39

AF002016 acyl CoA oxidase ‘Kurokawa Amakuri Nankin’ 9

PP36 AF274589 cytochrome b5 reductase C. maxima ‘Big Max’ **

pAPX AB070626 peroxisomal ascorbate peroxidase ‘Kurokawa Amakuri Nankin’ 33

CM-ACS3 AB038559 ACC synthase C. maxima 43

CmATS AB049135 acyl-(acyl-carrier protein); acyltransferase C. moschata ‘Shirogikuza’ **

Y00771 glycerol-3-phosphate acyltransferase transit peptide

C. moschata ‘Shirakikuza’ 17

AB002695 aspartic endopeptidase C. pepo 14

PS-1 AF284038 phloem serpin C. maxima 46

SLW AF170086, AF170087

silverleaf whitefly-induced protein (multiple genes)

C. pepo zucchini ‘Chefini’ 41

aprX Y17192 anionic peroxidase C. pepo zucchini ‘Black Beauty’ 3

cpCPK1 U90262 calcium-dependent calmodulin-independent protein kinase

C. pepo zucchini 6

PP16 AF079170, AF079171

mRNA movement protein; phloem transport (multiple alleles)

C. maxima ‘Big Max’ 44

AOBP D45066 transcription factor binding to ascorbate oxidase

C. maxima 22

accW D01032 auxin-induced 1-aminocyclopropane-1-carboxylate synthase

C. maxima ‘Ebisu’ 32

U37774 auxin-induced 1-aminocyclopropane-1-carboxylic acid synthase

C. maxima 31

ACC1 M58323 1-aminocyclopropane-1-carboxylate synthase

C. pepo 34

Page 19: Gene List for Cucurbita species, 2004 - Cucurbit Breedingcuke.hort.ncsu.edu/cgc/cgcgenes/gene04squash.pdf · Gene List for Cucurbita species, 2004 Harry S. Paris A.R.O., Newe Ya’ar

ACC1A, ACC1B

M61195 1-aminocyclopropane-1-carboxylate synthase (2 genes, tightly linked)

C. pepo zucchini 15

PHP-1 D86306 proton-translocating inorganic pyrophosphatase

C. moschata crookneck **

PP1 U66277 phloem filament protein C. maxima ‘Big Max’ 4

pfiAF4 X81647 trypsin inhibitor C. maxima ‘Supermarket Hybrid’ 30

pfiBM7 X81447 chymotrypsin inhibitor C. maxima ‘Supermarket Hybrid’ 30

M15265 phytochrome C. pepo zucchini ‘Black Beauty’ 35

NADH M33154 nitrate reductase C. maxima 5

M36407 11S globulin beta-subunit ‘Kurokawa Amakuri Nankin’ 11

AF206895 18S ribosomal RNA C. pepo **

AF479108 26S ribosomal RNA C. pepo 38

* Gene symbols were assigned by the researchers isolating the gene; they have no correspondence to the official Cucurbita gene symbols. **Unpublished: Genes can be submitted directly to Genbank, wthout being published in a journal. *** ‘Kurokawa Amakuri Nankin’ was identified only as “Cucurbita sp.” Literature Cited in Table 4 1. Aoki, K., F. Kragler, B. Xoconostle-Cazares and W.J. Lucas. 2002. A subclass of plant heat shock cognate 70

chaperones carries a motif that facilitates trafficking through plasmodesmata. Proc. Natl. Acad. Sci. U.S.A. 99: 16342-16347.

2. Bostwick, D.E. and G.A. Thompson. 1993. Nucleotide sequence of a pumpkin phloem lectin cDNA. Plant Physiol. 102: 693-694.

3. Carpin, S., M. Crevecoeur, H. Greppin, and C. Penel. 1999. Molecular cloning and tissue-specific expression of an anionic peroxidase in zucchini. Plant Physiol. 120: 799-810.

4. Clark, A.M., K.R. Jacobsen, D.E. Bostwick, J.M. Dannenhoffer, M.I. Skaggs and G.A. Thompson. 1997. Molecular characterization of a phloem-specific gene encoding the filament protein, phloem protein 1 (PP1), from Cucurbita maxima. Plant J. 12: 49-61.

5. Crawford, N.M., W.H. Campbell and R. Davis. 1986. Nitrate reductase from squash: cDNA cloning and nitrate regulation. Proc. Natl. Acad. Sci. U.S.A. 83: 8073-8076.

6. Ellard-Ivey, M., R.B. Hopkins, T.J White, and T.L. Lomax. 1999. Cloning, expression and N-terminal myristoylation of CpCPK1, a calcium-dependent protein kinase from zucchini (Cucurbita pepo L.). Plant Mol. Biol. 39: 199-208.

7. Esaka, M., T. Hattori, K. Fujisawa, S. Sakajo, and T. Asahi. 1990. Molecular cloning and nucleotide sequence of full-length cDNA for ascorbate oxidase from cultured pumpkin cells. Eur. J. Biochem. 191: 537-541.

8. Hara-Hishimura, I., Y. Takeuchi, K. Inoue and M. Nishimura. 1993. Vesicle transport and processing of the precursor to 2S albumin in pumpkin. Plant J. 4: 793-800.

9. Hayashi, H., L. De Bellis, K. Yamaguchi, A. Kato, M. Hayashi, and M. Nishimura. 1998. Molecular characterization of a glyoxysomal long chain acyl-CoA oxidase that is synthesized as a precursor of higher molecular mass in pumpkin. J. Biol. Chem. 273: 8301-8307.

10. Hayashi, H., L. De Bellis, A. Alpi and M. Nishimura. 1995. Cytosolic aconitase participates in the glyoxylate cycle in etiolated pumpkin cotyledons. Plant Cell Physiol. 36: 669-680.

11. Hayashi, M., H. Mori, M. Nishimura, T. Akazawa and I. Hara-Nishimura. 1988. Nucleotide sequence of cloned cDNA coding for pumpkin 11-S globulin beta subunit. Eur. J. Biochem. 172: 627-632.

12. Hayashi, M., R. Tsugeki, M. Kondo, H. Mori and M. Nishimura. 1996. Pumpkin hydroxypyruvate reductases with and without a putative C-terminal signal for targeting to microbodies may be produced by alternative splicing. Plant Mol. Biol. 30: 183-189

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13. Helliwell, C.A., M.R. Olive, L. Gebbie, R. Forster, W.J. Peacock, and E.S. Dennis. 2000. Isolation of an ent-kaurene oxidase cDNA from Cucurbita maxima. Aust. J. Plant Physiol. 27: 1141-1149.

14. Hiraiwa, N., M. Kondo, M. Nishimura, and I. Hara-Nishimura. 1997. An aspartic endopeptidase is involved in the breakdown of propeptides of storage proteins in protein-storage vacuoles of plants. Eur. J. Biochem. 246: 133-141.

15. Huang, P.L., J.E. Parks, W.H. Rottmann, and A. Theologis. 1991. Two genes encoding 1-aminocyclopropane-1-carboxylate synthase in zucchini (Cucurbita pepo) are clustered and similar but differentially regulated. Proc. Natl. Acad. Sci. U.S.A. 88: 7021-7025.

16. Inoue, K., Y. Takeuchi, M. Nishimura and I. Hara-Nishimura. 1995. Characterization of two integral membrane proteins located in the protein bodies of pumpkin seeds. Plant Mol. Biol. 28: 1089-1101.

17. Ishizaki, O., I. Nishida, K. Agata, G. Eguchi and N. Murata. 1988. Cloning and nucleotide sequence of cDNA for the plastid glycerol-3-phosphate acyltransferase from squash. FEBS Lett. 238: 424-430.

18. Kato, A., M. Hayashi, H. Mori, and M. Nishimura. 1995. Molecular characterization of a glyoxysomal citrate synthase that is synthesized as a precursor of higher molecular mass in pumpkin. Plant Mol. Biol. 27: 377-390.

19. Kato, A., M. Hayashi, Y. Takeuchi and M. Nishimura. 1996. cDNA cloning and expression of a gene for 3-ketoacyl-CoA thiolase in pumpkin cotyledons. Plant Mol. Biol. 31: 843-852.

20. Kim, M.G., K.O Lee, N.E. Cheong, Y.O. Choi, J.H. Jeong, M.J. Cho, S.C. Kim, and S.Y. Lee. 1999. Molecular cloning and characterization of a class III chitinase in pumpkin leaves, which strongly binds to regenerated chitin affinity gel. Plant Sci. 147: 157-163.

21. Kisu, Y., Y. Harada, M. Goto and M. Esaka. 1997.Cloning of the pumpkin ascorbate oxidase gene and analysis of a cis-acting region involved in induction by auxin. Plant Cell Physiol. 38: 631-637.

22. Kisu, Y., T. Ono, N. Shimofurutani, M. Suzuki, and M. Esaka. 1998. Characterization and expression of a new class of zinc finger protein that binds to silencer region of ascorbate oxidase gene. Plant Cell Physiol. 39: 1054-1064.

23. Klebl, F., M. Wolf, and N. Sauer. 2003. A defect in the yeast plasma membrane urea transporter Dur3p is complemented by CpNIP1, a Nod26-like protein from zucchini (Cucurbita pepo L.), and by Arabidopsis thaliana delta-TIP or gamma-TIP. FEBS Lett. 547: 69-74.

24. Lange, T. 1997. Cloning gibberellin dioxygenase genes from pumpkin endosperm by heterologous expression of enzyme activities in Escherichia coli. Proc. Natl. Acad. Sci. U.S.A. 94: 6553-6558.

25. Lange, T., P. Hedden, and J.E. Graebe. 1994. Expression cloning of a gibberellin 20-oxidase, a multifunctional enzyme involved in gibberellin biosynthesis. Proc. Natl. Acad. Sci. U.S.A. 91: 8552-8556.

26. Lange, T., S. Robatzek, and A. Frisse. 1997. Cloning and expression of a gibberellin 2 beta,3 beta-hydroxylase cDNA from pumpkin endosperm. Plant Cell 9: 1459-1467.

27. Mano, S., M. Hayashi, M. Kondo, and M. Nishimura. 1996. cDNA cloning and expression of a gene for isocitrate lyase in pumpkin cotyledons. Plant Cell Physiol. 37: 941-948.

28. Mano, S., K. Yamaguchi, M. Hayashi, and M. Nishimura. 1997. Stromal and thylakoid-bound ascorbate peroxidases are produced by alternative splicing in pumpkin. FEBS Lett. 413: 21-26.

29. Mori, H., Y. Takeda-Yoshikawa, I. Hara-Nishimura, and M. Nishimura. 1991. Pumpkin malate synthase. Cloning and sequencing of the cDNA and northern blot analysis. Eur. J. Biochem. 197: 331-336.

30. Murray, C. and J.T. Christeller. 1995. Purification of a trypsin inhibitor (PFTI) from pumpkin fruit phloem exudate and isolation of putative trypsin and chymotrypsin inhibitor cDNA clones. Biol. Chem. Hoppe-Seyler 376: 281-287.

31. Nakagawa, N., Y. Kamiya, and H. Imaseki. 1995. Nucleotide sequence of an auxin-regulated 1-aminocyclopropane-1-carboxylic acid synthase gene (Accession No. U37774) from Cucurbita maxima Duch. (PGR95-110). Plant Physiol. 109: 1499.

32. Nakajima, N., H. Mori, K. Yamazaki, and H. Imaseki. 1990. Molecular cloning and sequence of a complementary DNA encoding 1-aminocyclopropane-1-carboxylate synthase induced by tissue wounding. Plant Cell Physiol. 31: 1021-1029.

33. Nito, K., K. Yamaguchi, M. Kondo, M. Hayashi, and M. Nishimura. 2001. Pumpkin peroxisomal ascorbate peroxidase is localized on peroxisomal membranes and unknown membranous structures. Plant Cell Physiol. 42: 20-27.

34. Sato, T., P.W. Oeller, and A. Theologis. 1991. The 1-aminocyclopropane-1-carboxylate synthase of Cucurbita. Purification, properties, expression in Escherichia coli, and primary structure determination by DNA sequence analysis. J. Biol. Chem. 266: 3752-3759.

35. Sharrock, R.A., J.L. Lissemore, and P.H. Quail. 1986. Nucleotide and amino acid sequence of a Cucurbita phytochrome cDNA clone: identification of conserved features by comparison with Avena phytochrome. Gene 47: 287-295.

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36. Shimada, T., M. Kuroyanagi, M. Nishimura, and I. Hara-Nishimura. 1997. A pumpkin 72-kDa membrane protein of precursor-accumulating vesicles has characteristics of a vacuolar sorting receptor. Plant Cell Physiol. 38: 1414-1420.

37. Smith, M.W., S. Yamaguchi, T. Ait-Ali, and Y. Kamiya. 1998. The first step of gibberellin biosynthesis in pumpkin is catalyzed by at least two copalyl diphosphate synthases encoded by differentially regulated genes. Plant Physiol. 118: 1411-1419.

38. Soltis, D.E., A.E. Senters, M.J. Zanis, S. Kim, J.D. Thompson, P.S. Soltis, L.P. Ronse de Craene, P.K. Endress, and J.S. Farris. 2003. Gunnerales are sister to other core eudicots: implications for the evolution of pentamery. Am. J. Bot. 90: 461-470.

39. Tsugeki, R., I. Hara-Nishimura, H. Mori, and M. Nishimura. 1993. Cloning and sequencing of cDNA for glycolate oxidase from pumpkin cotyledons and northern blot analysis. Plant Cell Physiol. 34: 51-57

40. Tsugeki, R., H. Mori, and M. Nishimura. 1992. Purification, cDNA cloning and Northern-blot analysis of mitochondrial chaperonin 60 from pumpkin cotyledons. Eur. J. Biochem. 209: 453-458.

41. van de Ven, W.T., C.S. LeVesque, T.M. Perring, and L.L. Walling. 2000. Local and systemic changes in squash gene expression in response to silverleaf whitefly feeding. Plant Cell 12: 1409-1423.

42. Wang, M.B., D. Boulter, and J.A. Gatehouse. 1994. Characterization and sequencing of cDNA clone encoding the phloem protein PP2 of Cucurbita pepo. Plant Mol. Biol. 24: 159-170.

43. Watanabe, T., H. Fujita, and S. Sakai. 2001. Effects of jasmonic acid and ethylene on the expression of three genes for wound-inducible 1-aminocyclopropane-1-carboxylate synthase in winter squash (Cucurbita maxima). Plant Sci. 161: 67-75.

44. Xoconostle-Cazares, B., Y. Xiang, R. Ruiz-Medrano, H.L. Wang, J. Monzer, B.C. Yoo, K.C. McFarland, V.R. Franceschi, and W.J. Lucas. 1999. Plant paralog to viral movement protein that potentiates transport of mRNA into the phloem. Science 283: 94-98.

45. Yamaguchi, K., M. Hayashi, and M. Nishimura. 1996. cDNA cloning of thylakoid-bound ascorbate peroxidase in pumpkin and its characterization. Plant Cell Physiol. 37: 405-409.

46. Yoo, B.C., K. Aoki, Y. Xiang, L.R. Campbell, R.J. Hull, B. Xoconostle-Cazares, J. Monzer, J.Y. Lee, D.E. Ullman, and W.J. Lucas. 2000. Characterization of Cucurbita maxima phloem serpin-1 (CmPS-1). A developmentally regulated elastase inhibitor. J. Biol. Chem. 275: 35122-35128.

47. Yoo, B.C., J.Y. Lee, and W.J. Lucas. 2002. Analysis of the complexity of protein kinases within the phloem sieve tube system. Characterization of Cucurbita maxima calmodulin-like domain protein kinase 1. J. Biol. Chem. 277: 15325-15332.